The history of freshwater research in the Philippines with notes on its origins in the University of Santo Tomas and present-day contributions

2017·1 MB·

by  Briones, Rey Donne S Papa And Jonathan Carlo A

REVIEW PAPER | Philippine Journal of Systematic Biology
The History of Freshwater Research in the Philippines with
Notes on its Origins in the University of Santo Tomas and
Present-Day Contributions
Rey Donne S. Papa1,2,3* and Jonathan Carlo A. Briones1
ABSTRACT

The study of freshwater ecosystems in the country has long had the reputation of being
fragmentary and inconsistent, especially when compared to its marine counterparts. Other
scientists have in fact noted that many studies that have been conducted on Philippine
freshwaters have not gone beyond the “age of exploration”, which resulted to it being
poorly represented in the scientific literature. This scenario has been consistent for both
lotic and lentic habitats, even for those considered as major river and lake ecosystems,
including those found in key biodiversity or known protected areas. This has gone on for
decades in spite of the obvious need for invigorated and scientifically-driven approaches to
study and manage freshwater ecosystems throughout the country, especially since

freshwater ecosystems, including its flora and fauna, are under increasing threat from both
KEY WORDS :
natural and man-made environmental stressors, including intensive aquaculture, leading to

eutrophication, the introduction of non-native species, as well as climate change. This is
Wallacea Expedition
further aggravated by the low number of experts in various fields of basic and applied
Tropical Lakes
History of Science freshwater biology including taxonomists, limnologists, restoration ecologists etc. to deal
Freshwater Zooplankton with various problems and challenges in the Philippine setting. This paper presents the
Invasive species background, history and origins of freshwater biology research in the Philippines and the
Limnology contributions of the University of Santo Tomas from 2002 to present.
INTRODUCTION freshwater ecosystems to some extent are Anacleto del
Rosario and Antonio Luna, who, in their capacity as
The University of Santo Tomas, being the oldest existing scientists of the Laboratorio Municipal de la Ciudad de
institution of higher learning in the Philippines, has had a Manila, separately conducted biological and chemical
significant role in the development of science in the country studies of water from the Pasig River and medicinal mineral
(Papa, 2013). In the fields of natural history and biology its springs in Luzon (Reyes, 2014). Line drawings of
Museum of Arts and Science served as the earliest known, microscopic specimens observed by Anacleto del Rosario
scientifically-catalogued repository of Philippine fauna from the Pasig River clearly depicts several species of
collected from the 1800’s through the efforts of Rev. Fr. Casto bacteria, microalgae (phytoplankton), rotifers and
de Elera, OP - second director of the UST Museum. The protozoans. These line drawings, which now form part of
publication of a three-volume book on the fauna of the the collections of the Archives of the University of Santo
Philippines by Fr. de Elera provided one of the earliest Tomas (AUST), may be considered as some of the earliest
accounts of Philippine biodiversity (De Elera, 1895). Though documentations of freshwater biodiversity in the country
not as popular as contributions to other fields of science (such (Papa, 2015). Both del Rosario and Luna studied
as chemistry, pharmacy and medicine), Thomasian alumni Pharmacy in the university. Del Rosario finished his
have also conducted freshwater biology research during the licentiate and doctorate in Pharmacy in UST where he was
latter part of the 19th century. Two notable alumni who studied also later appointed Professor of Pharmacy while Luna
began his pharmacy studies in Santo Tomas but finished
1Department of Biological Sciences, College of Science his doctorate degree in the Universidad Central de Madrid
2The Graduate School 3Research Center for the Natural and Applied and took post-doctoral training in the Pasteur Institute
Sciences, University of Santo Tomas, Manila, Philippines 1015 (Paris, France) prior to serving the Laboratorio Municipal.
Corresponding Author: [email protected] Luna, of course, later became more well known for his role
Date Submitted: 03 September 2016
as a general in the Philippine revolutionary army (Papa,
Date Accepted: 01 January 2017
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2013). Though technically graduates of pharmacy, these government scientists had also included some notes on
individuals performed scientific duties other than those done major freshwater ecosystems in the country as part of the
by pharmacists as we know today; this was due to the lack of US government’s effort to document the natural resources of
any other related science course being offered in the country. their newly acquired colony (Pratt, 1916). Another notable
Del Rosario is credited as the “Father of Philippine Laboratory publication by Pratt documented the impacts of the 1911
Science” and the first Filipino chemist, while Luna, after his eruption of Taal Volcano, which also noted the eruption’s
and post-doctoral training, was regarded as the first published impacts on Lake Bombon, now more popularly known as
bacteriologist in the country (Papa, 2013). Lake Taal (Pratt, 1911).

This paper outlines the development and the significant GOLDEN AGE
contributions of the University of Santo Tomas, particularly its
Research Center for the Natural and Applied Sciences, to the German developmental zoologist Richard Woltereck of the
field of freshwater biology and taxonomy during the past University of Leipzig led the most comprehensive biodiversity
twelve years, where it has conducted research projects within survey of major lake ecosystems in the country when he
the general realm of freshwater biology. This has led to the headed the U.S. Rockefeller Foundation-funded Wallacea
group receiving funding and awards from national and (Woltereck-Tressler) Expedition in March 1932, which
international award / grant-giving bodies and the publication formed part of sixteen-month attempt to study ecologically-
of no less than 30 peer-reviewed papers. This is especially distinct Southeast Asian lakes. For the longest time, this had
significant since prior to this, the university’s research agenda been considered as the most comprehensive documentation
did not even include biodiversity and ecology priority areas. of freshwater flora and fauna in the country with descriptions
Difficulties that have also evolved due to the location of the of limnological characteristics for many of the lakes they had
university in the heart of Manila did not hinder its visited. This expedition resulted to several landmark
development. Through the efforts of its faculty and students, publications in journals such as the Internationale Revue der
freshwater scientists from the University of Santo Tomas have Gesamten Hydrobiologie und Hydrogeographie and
been able to continue and build on the legacy began by its Zoologischer Anzeiger (Woltereck et al. , 1941). As for
illustrious alumni - Anacleto del Rosario and Antonio Luna. studies on freshwater fishes, the American Albert William T.
Herre (Director of the Fisheries Division in the Philippine
AGE OF EXPLORATION Bureau of Science), contributed the most significant scientific
information through his 216 publications written over the
Though such pioneering studies on the biology and chemistry span of his entire career, which included novel descriptions
of Philippine freshwaters have been contributed by scientists of many freshwater fishes native or endemic to the country
who, at one point in their careers had been associated with (Herre, 1953).
Santo Tomas, these never got the same attention as those
from German naturalist Feodor Jagor, who has written Even though scientific literature on Philippine freshwaters
descriptions of Philippine lakes he had visited during his during this time was dominated by foreign authors, there
expeditions, as documented in his book “Travels in the were two Filipinos who contributed to the development of
Philippines” (1875), which also included collections of its flora freshwater biology in the country. The first Filipino to have
and fauna (Buñag, 1934). Towards the end of the 19th published papers on the topic is considered the “Dean of
Century, another major publication that included listings and Philippine Fisheries” - Deogracias V. Villadolid, who taught at
descriptions of freshwater ecosystems came in the form of the University of the Philippines College of Agriculture (now
Jose Montero y Vidal’s “El Archipelago Filipino” (1896 & the University of the Philippines Los Baños) after finishing
1909) (Vidal, 1909). his Ph.D. in Stanford University. His earliest works included
papers on the fisheries of Lake Taal and Laguna de Bay, as
The end of the Spanish colonization of the Philippine well as Pansipit River (Villadolid and Del Rosario, 1930;
archipelago and the start of the American regime opened up Villadolid, 1932a, b, c; Villadolid, 1937). He later became
more opportunities for other scientists to study Philippine director of the Bureau of Fisheries where he was
freshwater ecosystems. In 1916, Wallace Pratt published a instrumental to the introduction of tilapia aquaculture in the
paper entitled “Philippine Lakes”, in the Philippine Journal of Philippines by the 1950’s. He was also influential to the
Science, which may be considered as the first listing of establishment of the UP College of Agriculture Limnological
Philippine lakes published in a scientific journal (Pratt, 1916). Research and Aquatic Resources Management Center,
His involvement with lake research came courtesy of his which was established through the proclamation of American
appointment in the Division of Mines in the Philippine Bureau Governor General Henry Stimson in 1928 upon the initiative
of Science. Prior to this, several geological studies by US of Leopoldo Uichanco (Dean of the UP College of
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Agriculture) and Albert William T. Herre (Labatos, conducted by Rafael D. Guerrero III on tilapia sex reversal
unpublished). The other Filipino to have made a significant are also considered among the most significant outputs on
contribution to the study of Philippine freshwater ecosystems applied researches in freshwater biology in the country (Lee-
was Daniel M. Buñag, also of the University of the Philippines Chua, 2000). Later, as director of the Philippine Council for
College of Agriculture, who wrote “A supplement to our Aquatic and Marine Research and Development, he
knowledge of two Philippine lakes” (Buñag, 1934) and co- published several papers on the conservation and proper
authored “Die Seen und Inseln der "Wallacea"-Zwishenregion utilization of lake resources (Guerrero III, 1991; Guerrero III,
und ihre endemische Tierwelt. Zweiter Teil: Inseln und Seen 2001, 2002). Of the studies mentioned above, the works of
der Philippinen” with R. Woltereck and W. Tressler (Woltereck William Lewis in Lakes Mainit and Lanao were some of the
et al. , 1941). The achievements during the early part of the first detailed observations on the limnology and ecology of
20th Century in freshwater biology may very well be tropical lakes that have been published in journals such as
considered as its “Golden Age”, when the scientific Limnology and Oceanography, Ecological Monographs and
community was introduced to the unique characteristics and the International Review of Hydrobiology.
biodiversity of freshwater habitats in the Philippines.
The Southeast Asian Fisheries Development Center
FILIPINIZATION (SEAFDEC), particularly its Binangonan Freshwater Station
in Laguna de Bay, also published important research papers
Sadly, the events of the Second World War halted any major on freshwater organisms, though many papers have dealt
developments. The subsequent granting of Philippine with aquaculture applications, others focused on the
independence in 1946, failed to revive interest in studying occurrence of harmful algal blooms, particularly in Laguna de
freshwater ecosystems in the country to similar levels it had Bay (Baldia et al. , 1986; Platon, 2001; Baldia et al. , 2003;
prior to the start of the war. The focus of most Filipino Baldia et al. , 2007). By the 1990’s, the freshwater biology
researchers shifted to the utilization of freshwater resources studies in the Philippines began to look into the negative
for food, out of necessity, which led to the introduction of impacts of intensive and irresponsible aquaculture in the
aquaculture in many Philippine lakes. Throughout the 1950’s Philippines. Results generated during this time have noted
to the 1960’s, efforts to study freshwater bodies as significant reductions in the water quality of lakes such as
ecosystems have almost been relegated to the sidelines, and Laguna de Bay, Lake Sampaloc and Lake Taal. This has
were not in the same level before the war. As a result, novel prompted the development of laws and policies related to
scientific information on Philippine freshwater ecosystems in lake zoning, the determination of the carrying capacity of
the form of peer-reviewed publications, went down lakes, and routine water quality monitoring (Pantastico et al. ,
significantly during this time (Papa and Mamaril Sr., 2011). 1986; Santiago et al. , 1989; Zafaralla et al. , 1989; Zafaralla
The succeeding three decades (1970’s to 1990’s) saw slight et al. , 1992; Santiago and Arcilla, 1993; Pctt, 1994; Lasco
improvement, which resulted to several international and and Espaldon, 2005; Baldia et al. , 2007; Schiemer et al. ,
national publications. This was due to the increased 2008; TVPL-PAMB, 2009). However, the combined
international exposure of some Filipinos who had been given detrimental effects of intensive aquaculture and the problems
the chance to undergo further training in foreign laboratories. that arose from a ballooning human population who settled
Significant outputs included papers on the limnology and previously unoccupied lakeshore or riverside areas have
plankton ecology of Lakes Mainit and Lanao by the American contributed to the rapid decline of freshwater ecosystems,
limnologist William Lewis Jr. (Lewis Jr., 1973b; Lewis Jr., particularly those near urban areas and in water-bodies
1973a, 1974; Lewis Jr., 1975, 1979; Babin et al. , 2005), throughout the Metropolis. This problem was compounded
updates on the taxonomy of Philippine freshwater by the practice of majority of Filipino freshwater scientists to
zooplankton by Augustus C. Mamaril (who studied under the either not publish or merely being contented with
renowned zooplankton taxonomist, C. H. Fernando of the publications in gray literature. This has been previously
University of Waterloo) (Mamaril Sr. and Fernando, 1978; Lai noted for studies in Philippine marine ecosystems
et al. , 1979; Mamaril Sr., 1986), phytoplankton by Macrina (Lacanilao, 1997), and Lake Taal from the 1960’s to the
Tamayo-Zafaralla and Milagrosa Martinez-Goss (Martinez, 1990’s (Papa and Mamaril Sr., 2011), but was also the case
1978; Zafaralla and Orozco, 1989; Zafaralla, 1990; Zafaralla for the other freshwater ecosystems that had been studied.
et al. , 1990; Zafaralla, 1998), littoral fishes and gastropods in
Lake Taal by Roberto C. Pagulayan (Pagulayan and Remigio, Since the start of Filipino involvement in this scientific field,
1992/3; Cukingan and Pagulayan, 1995-1996; Pagulayan et biologists from the University of the Philippines and / the
al. , 1997), as well as a textbook / laboratory manual on Bureau of Fisheries and Aquatic Resources / the Southeast
limnology designed for the Philippine setting by Ruben Umaly Asian Fisheries Development Center / Laguna Lake
and Ma. Lourdes Cuvin (Umaly and Cuvin, 1988). The studies Development Authority have been at the forefront, trying to
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sustain the fledgling field of freshwater science in the country, freshwater biology researches being conducted by the
which, when compared to other more popular fields of university. This may be due to the fact that none of those
specialization in the natural and applied sciences, had who taught freshwater biology and other related biology
received far little attention, students and funding. Many of courses were actually affiliated with the Research Center for
these researches have been funded by the Philippine Council the Natural Sciences – the university unit devoted to the
for Aquatic and Marine Research and Development conduct of research projects in the sciences.
(PCAMRD) locally and internationally by funding agencies
such as the SOGREAH (Société Grenobloise d’Etudes et d’ All this changed when in 2002, Dr. Susana F. Baldia, who
Applications Hydrauliques) and the Danish International was formerly affiliated with the SEAFDEC Binangonan
Development Agency (DANIDA). Much attention has been Freshwater Station, joined the graduate faculty in Santo
focused on studying Laguna de Bay, making it the most Tomas, where she taught courses in Freshwater Biology,
widely studied Philippine lake, to date. Advanced Ecology and Phycology. Apart from teaching in
the Graduate School, she also conducted researches on
FRESHWATER BIOLOGY IN SANTO TOMAS 1: BIRTHING PAINS freshwater phytoplankton in the UST Research Center for
the Natural Sciences (RCNS), where she pioneered
It was against this backdrop that, after the pioneering efforts research on freshwater organisms and later began to teach
of del Rosario and Luna more than a century before, that UST in the undergraduate level as well. She was later joined by
scientists began to make contributions to freshwater biology. Alicia Ely J. Pagulayan, a faculty member of the College of
No doubt, through the influence of professors from the Science, who conducted research on the developmental
University of the Philippines, but similarly from scientists biology of the Lake Taal endemic freshwater sardine,
affiliated with other local and international institutes and Sardinella tawilis. In 2006, the author (R.D.S. Papa) also
universities, who had the chance to interact and influence a joined the UST-RCNS after completing his M.Sc. thesis
new breed of eager-to-learn Thomasian biologists. These under the supervision of Dr. R.C. Pagulayan and A.E.J.
individuals saw the need to increase awareness and scientific Pagulayan where he worked on the zooplankton diet of the
knowledge on a vital, yet understudied component of our Sardinella tawilis in Lake Taal. His RCNS research project
environment. The efforts come during a time when freshwater focused on ecological studies on freshwater zooplankton
resources in the country are faced with numerous threats and from different lakes which he was doing simultaneously with
challenges, from both man-made and natural causes, that are his academic requirements for his Ph.D. These
being aggravated by a changing climate. developments in the university have opened opportunities for
undergraduate and eventually graduate students to conduct
The initial exposure of Thomasians to the field of freshwater freshwater biology researches in at least three fields:
biology was due to the influence of faculty members of the phytoplankton, zooplankton and fish development.
Graduate School, as the undergraduate program in the
College of Science did not have a freshwater biology course. Lake Taal became the initial focus of UST researchers,
These faculty members were usually part-timers with regular which proved to be an enduring and fruitful endeavor. The
teaching or research appointments in other universities or initial studies conducted by A.E.J. Pagulayan and R.D.S.
government research institutions. During the 1970’s and Papa for their master’s theses were followed by several
1980’s, Dr. Ruben Umaly of UP Diliman, author of the undergraduate theses. In 2006, a study by undergraduate
laboratory and field guide in limnology (National Bookstore, students of R.D.S. Papa and molecular biologist J.R. Castillo
1988), taught genetics and radiation biology in the UST on the morphological and molecular identification of the
Graduate School. Several theses conducted by UST graduate ectoparasitic isopod Corallana grandiventra (locally known
students in the 1970’s also dealt with freshwater biology. This as timud) was published in The Philippine BIOTA Journal,
included the bio-ecology of the freshwater gastropod making it the first published paper on Lake Taal fauna based
Idiopoma angularis in Laguna de Bay (Castro, 1977) as well on an undergraduate thesis from the university(Adorador et
as a survey of monogenean parasites of fishes in Laguna de al. , 2006) . This paper was able to correct the previous
Bay (Guzman, 1974). By the 1990’s, the graduate course in misidentification of the Lake Taal isopod which appeared in
Freshwater Biology had been taught by Dr. Abercio V. Rotor, several reports by the local Bureau of Fisheries and Aquatic
a former student of Dr. Villadolid, who also taught Advanced Resources office. Two years later, the first ISI publication
Ecology and several botany courses. He was later joined by based on a UST Graduate School master’s thesis in
Dr. Roberto C. Pagulayan, who taught genetics and ecology freshwater biology came out in the journal Zoological
from the late 1990’s to the early 2000’s. Though freshwater Studies. The study documented the feeding ecology of the
biology was regularly taught in the Graduate School, there Sardinella tawilis in Lake Taal (Papa et al. , 2008a). Apart
had been no significant development in the publication of from these major developments, active participation of
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Thomasian researchers in national and international in foreign laboratories located in institutions such as the
conferences and training seminars “introduced” UST’s efforts National Taiwan University (Taiwan), Jinan University
to undertake research projects on freshwater biology to the (China), Polish Academy of Science (Poland), Kyoto
scientific community. University (Japan), National Institute for Agronomy Research
(France), University of Shiga Prefecture (Japan), and
FRESHWATER BIOLOGY IN SANTO TOMAS 2: MILESTONES National University of Singapore (Singapore). This led to the
increased number of quality publications in freshwater
The developments from the years 2002 to 2008 laid the biology produced by the University of Santo Tomas, and also
foundations for the increased number of research staff and enabled it to apply for grants to support its researches. The
students from the University of Santo Tomas involved in following sections highlight the major outputs of the research
freshwater biology research. In 2011, the increased number group.
of students engaged in freshwater researches led to the
formation of the Zooplankton Ecology, Systematics and Zooplankton Systematics
Limnology research group in the Research Center for the Perhaps, the most significant contributions of the university
Natural and Applied Sciences. Popularly known through its are in the field of freshwater zooplankton systematics. There
acronym ZESL, the group conducted research projects which had been very few publications that had come out on
involved graduate and undergraduate students, and many of Philippine freshwater zooplankton to build up on the work of
these were under the framework of national and international A. C. Mamaril from the 1970’s to the 1990’s (Tuyor and
collaborative research projects. International collaborations Segers, 1999; Tuyor and Baay, 2001). The advances in the
opened up venues for students to learn specific aspects of taxonomy and systematics of freshwater zooplankton
freshwater biology researches that they would not have worldwide, together with the changes that have occurred in
otherwise been given the opportunity to learn given the terms of the utilization of freshwater resources makes it
limitations in the country. Foreign collaborators have been necessary to conduct a re-evaluation of the state of
invited to deliver lectures, conduct training seminars and also freshwater zooplankton diversity. In order to do this, faculty
teach graduate-level courses. More importantly, students and students from the University of Santo Tomas conducted
were given the opportunity to conduct part of their researches sampling expeditions to different parts of the country to 1)
Figure 1. Sampling localities of zooplankton samples deposited in the Zooplankton Reference
Collection (ZRC) of the Research Center for the Natural and Applied Sciences – University of
Santo Tomas
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revisit former sampling localities where zooplankton have species that were displaced by A. dorsalis are the endemic
been previously recorded and 2) collect samples from areas Tropodiaptomus gigantoviger in Lake Lanao, the native T.
where no previous records have existed. Figure 1 shows the vicinus in Lake Buhi and the endemic Pseudodiaptomus
extent of sampling areas where zooplankton had been brehmi in Lake Naujan. This is especially significant for
collected from different parts of the country. During the first calanoid copepods, as there is usually just one calanoid
two years of the ZESL, a research project entitled copepod present for any given ecosystem. Notes on the
“Establishing a Zooplankton Reference Collection in the mode of dispersal and its successful invasion are available in
University of Santo Tomas” was approved and funded by the Papa et al. (2012) (Papa et al. , 2012a). In another group of
RCNAS, in order to come up with an organized system of copepods, the Order Cyclopoida, however, there have been
managing specimens and data gathered from the different no documented invasive species among the species
expeditions. The protocols established as a result of this identified from samples collected from 22 lakes found in 5
project were benchmarked from foreign institutions with major islands throughout the country (Papa and Holynska,
established zooplankton reference collections, and were 2013). In fact, a new species, Mesocyclops augusti Papa
modified to suit the local situation (Figure 2). and Holynska, 2013 was described from samples collected
from Lake Siloton (Mindanao Is.). The same paper also
contains a redescription of the endemic Mesocyclops
microlasius Kiefer 1981, where recent expeditions revealed it
was also present in Lake Paoay (Luzon Is.), making it the
northernmost limit of its distribution, to date. A total of 11
taxa of cyclopoid copepods were also identified from all the
samples collected from all 22 lakes, including
Thermocyclops taihokuensis (Harada, 1931), a new
Philippine record.

The other major contribution to freshwater zooplankton
systematics dealt with the taxonomy and distribution of
freshwater water fleas (Branchiopoda: Cladocera) in the
Philippines. The paper by Pascual et al. (Pascual et al. ,
2014), was able to identify a total of 16 species from four
cladoceran families (Bosminidae, Moinidae, Sididae,
Chydoridae) which were collected from 86 freshwater
Figure 1. The organization of samples in the University of Santo ecosystems found in six islands throughout the country. The
Tomas – Zooplankton Reference Collection A) part of the specimen 16 species which were identified is extremely low compared
cabinet showing unsorted samples B) sorted specimens C) slides to the 56 species previously recorded throughout the
and D) paratype of Mesocyclops augusti Papa and Holynska, 2013.
country, but included 78 new locality records. The study was
also able to note that most of the 56 species previously
An updated species list and distribution map for recorded from the country have now been relegated as
microcrustacean zooplankton is given in Figure 3. This synonyms, or have been misidentified.
includes several new locality and new species records for the
Philippines and one new species. Among the significant The major findings from all three studies (calanoid, cyclopoid
discoveries was the discovery of the presence of a and cladoceran taxonomy and distribution) were done using
Neotropical species Arctodiaptomus dorsalis (Calanoida: the standard methods used in zooplankton sampling,
Diaptomidae) in Philippine lake and river ecosystems (Papa processing and analysis that had been developed in the
et al. , 2012a; Metillo et al. , 2014; Rizo et al. , 2015). The university. This has enabled students and researchers to
occurrence of this species was discovered by accident, as the utilize previously collected samples for taxonomic analysis
original intention of the study was to try to increase the that has reduced the need to conduct repetitive sampling
number of known calanoid copepod species known from the expeditions, unless otherwise needed. As such, more efforts
Philippines. Instead, it led to the discovery of the presence of have been put into sampling sites where no previous
this non-indigenous zooplankton species in the Philippines. samples have been collected. At present, the University of
Unfortunately, previously recorded calanoid copepods were Santo Tomas is the only institution with a Zooplankton
not found in sites where A. dorsalis had been recorded. This Reference Collection in the Philippines.
confirms that A. dorsalis is highly invasive, and can displace
species previously inhabiting the ecosystem. Among the
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Figure 3. Distribution map of microcrustacean zooplankton based on published papers by researchers from the University of Santo Tomas
(Papa et al., 2012; Papa and Holynska, 2013; Pascual et al., 2014)
Ecology produce lipids, further studies by students of Dr. S. Baldia
In terms of freshwater ecology, most efforts have been site- led to several publications that dealt with the optimization of
specific, but ensured that sampling efforts were conducted for culture conditions of B. braunii isolates from Lake Paoay to
an adequate period of time. The first published studies on explore the possibility of using it for bio-fuel research, as well
freshwater ecology from the university were based on as a year-long sampling of phytoplankton in Lake Paoay to
undergraduate theses, that have been conducted in Lake look at the other phytoplankton species in the area
Paoay (Aquino et al. , 2008) and the Pasig River (Lazo et al. , (Villaroman et al. , 2010; Baldia et al. , 2011).
2009). Though the study in Lake Paoay originally focused on
zooplankton community dynamics, the discovery of a six- From 2008 to 2014, most papers published on freshwater
month continuous bloom of the colonial green algae, ecology that came from UST had been conducted in Lake
Botryococcus braunii led to a follow-up study that looked at Taal. These studies, which focused on 1) feeding ecology of
the negative impacts of the bloom to the zooplankton the Sardinella tawilis (Papa et al. , 2008a), 2) the spatio-
community (Papa et al. , 2008b). Since B. braunii is known to temporal variability and diel vertical migration in freshwater
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zooplankton (Papa and Zafaralla, 2011; Papa et al. , 2011; site for aquaculture in the 1970’s (Santiago et al. , 2001). By
Papa et al. , 2012b), as well as the 3) diet and distribution of the late 1980’s, however, floating cages for tilapia culture in
the endemic sea snake Hydrophis semperi (Garcia et al. , the lake expanded from 0.06 to 0.33 km2 of its total surface
2014). area, with at most 6,000 t of fish feeds used in the lake
annually (Santiago and Arcilla, 1993). The resulting increase
In 2011, Dr. Flor Lacanilao, former SEAFDEC Director and in allochthonous organic matter caused the worst
retired professor of marine science in the University of the occurrences of mass fish kills in Lake Sampaloc in the early
Philippines - Diliman made a survey of institutions with 1990’s, resulting in revenue loss and the risk of lake faunal
published researches on Laguna de Bay, Lake Taal and extinction. These events prompted the revision of
Bolinao (Pangasinan), which were among the most studied aquaculture and integrated water resources management
aquatic ecosystems in the country. His results showed that plans in the lake which resulted to the reduction of the
UST had the most number of published papers (ISI-indexed) allowable cage area to only 0.12 km2 which is approximately
on Lake Taal, followed closely by the University of the 10 % of the lake surface area (Cariño, 2003). These new
Philippines – Los Baños (Lacanilao, unpublished). provisions were geared towards lake rehabilitation and the
protection of local stakeholders thru responsible aquaculture
Another lake ecosystem that had been the focus of studies by practices. However, just recently, Global Nature Fund (GNF)
faculty and students of UST is Lake Sampaloc, which is proclaimed Lake Sampaloc as the “Threatened Lake of the
among many small lakes in the Philippines that have been Year 2014”. GNF cited the local government’s difficulty in
tapped as an economic resource, most notably as a pioneer implementing the imposed regulations due to inadequacy of
Table 1. List of parasites obtained from different host fishes and sampling locality
Classification Species Host Locality Remarks
ACANTHOCEPHALA
Polymorphida Bolbosoma sp. Oreochromis niloticus Lake Taal
Neoechinorhynchus
Neoechinorhynchida Carassius gibelio Lake Sampaloc All acanthocephalans
quinghaiensis
listed herein are first
Hypopthalmichthys
Laguna de Bay records in the
nobilis
Philippines (Briones et
Lake Sampaloc, al. in press)
Oreochromis niloticus
Laguna de bay
Rhadinorhynchus
Echinorhynchida Katsuwonis pelamis Batangas Sea
ganapatti
ARTHROPODA
Among the pioneering
studies which
confirmed the
Isopoda Corallana grandiventra Oreochromis niloticus Lake Taal
presence of Corallana
in Lake Taal
(Adorador et al. 2006)
Copepoda Lernaea sp. Oreochromis niloticus Lake Taal
PLATYHELMINTHES
Strigeidida Clinostomum sp. Clarias batrachus Lake Taal All platyhelminth
Euclinostomum sp. Clarias batrachus Lake Taal parasites listed herein
are among the few
Azygiida Erileptus sp. Glossogobius giuris Lake Taal
published works on
Plagiorchiida Orieantocreadium sp. Clarias batrachus Lake Taal fish parasites in Lake
Taal (Cauyan et al.
Opegaster sp. Clarias batrachus Lake Taal
2013)
Glossogobius giuris Lake Taal

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Philippine Journal of Systematic Biology | PAPA R.D.S & BRIONES J.C.A.: The History of Freshwater Research in the Philippines
Lumanlan-Mayo (Arthur and Lumanlan-Mayo, 1997)
manpower and funding, and highlighted the continuing
organized local reports into the Checklist of the parasites of
degradation of the Lake Sampaloc ecosystem. This issue has
fishes of the Philippines, which remains the most extensive
raised concerns regarding the current status and potential
reference for Philippine fish parasites to date. The report
threats to Lake Sampaloc’s biodiversity (Fund, 2014).
compiled records of fish parasites in the Philippines, and
However, to date, most of the published research done in the
amounted to 201 parasite species from 190 species of fish
lake has been related only to aquaculture implications
host examined. FishBase records (viewed November 2015)
(Santiago and Arcilla, 1993; Tan et al. , 1995; Santiago et al. ,
estimate a total of 3282 fish species in the Philippines to
2001; Cariño, 2003) with only a handful of studies on the lake’
date. Given that many parasites are potentially host-specific,
s actual biota (Quilang et al. , 2007; Santos et al. , 2010). As
we can only infer that there still remains a great deal of
a result, there is a gap on what is currently known regarding
taxonomic and basic survey work before the parasites of
the lake’s biodiversity. Establishment of baseline information
Philippine fishes have been thoroughly documented.
regarding the various biota of Lake Sampaloc is therefore

needed. Thomasian scientists have conducted biodiversity
Biodiversity-driven parasite research is still in the exploratory
surveys in many small lakes, and have placed particular focus
stage in the Philippine setting. The initiatives of students and
on the zooplankton, zoobenthos, and nekton of Lake
faculty of the University of Santo Tomas to lessen this gap
Sampaloc during their rapid surveys during 2012-2014. A
has been to focus on basic parasite survey work with
highlight to their work is the first attempt at the official listing of
implications to the systematics, biodiversity, and ecology of
fish (Briones et al. , 2016) and benthos from the lake. These
freshwater fish parasites in the Philippines. Highlights to their
findings have helped elucidate the present dominance of
work are first reports of Acanthocephala from Philippine fish:
many introduced biota from these lakes.
namely Neoechinorhynchus quinghaiensis,

Rhadinorhynchus ganapatti, and Bolbosoma sp. These three
The contribution of Thomasian freshwater biologists was not
new records encompass a third of all the known species of
confined to local ecosystems. Through collaborative
acanthocephalan parasites associated with Philippine fishes
agreements with scientists from the National Taiwan
(Briones et al. , 2015). Also, pioneering surveys on the
University and Kyoto University, a Thomasian was able to
lead the analysis of the long-term changes in the diet of the
Table 2. The occurrence records (non-native, native and endemic)
endemic goby Gymnogobius isaza in Lake Biwa. This was
of the different host fishes where the parasites have been obtained
done by analyzing the stomachs of archived specimens kept
in Kyoto University that had been collected over the span of Classification Species Occurrence
40 years (Briones et al., 2012).
CYPRINIFORMES

Fish Parasites Cyprinidae Carassius gibelio non-native
The Philippines has a rich history in fish parasite-related
Hypophthalmichthys nobilis non-native
research, many of which have pioneered the knowledge in
parasite biodiversity in tropical Asia. The monograph PERCIFORMES
Digenetic Trematodes of Philippine Fishes (Velasquez, 1975)
Channidae Channa striata non-native
is considered the first comprehensive monograph in this
region. Likewise, the extensive records of helminths of Cichlidae Oreochromis niloticus non-native
Philippine animals by Tubangui (Tubangui, 1933) together
Parachromis managuensis non-native
with other recent records, have contributed a great bulk in the
fish health literature in Southeast Asia by the end of the 20th Eleotridae Giuris margaritacea native
century. However, notwithstanding the great deal of research Gobiidae Glossogobius aureus native
already accomplished, a huge gap is still unknown for Terapontidae Leiopotherapon plumbeus endemic
Philippine fish parasite biodiversity in general. SILURIFORMES

Clariidae Clarias batrachus non-native
In the Philippines, studies on parasite hosts have been
Pangasianodon hy-
inclined towards aquaculture and aquarium fishes. This trend, Pangasiidae non-native
pophthalmus
unsurprisingly, is focused on detecting epizootics and
avoiding the negative impact of parasites in commercial HYBRID CICHLIDS
fisheries. Such an approach primarily employed parasite Oreochromis aureus
non-native
surveys from laboratory-based fish enclosures and fish Oreochromis mossambicus
market areas, making it difficult to infer specific locality Cichlosoma Amphilophus
non-native
records for parasites in Philippine inland waters. Arthur and Paraneetroplus
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Philippine Journal of Systematic Biology | PAPA R.D.S & BRIONES J.C.A.: The History of Freshwater Research in the Philippines
platyhelminth parasites of fish from Lake Taal have assessed the Asian Fisheries Society.
initial measures of parasite burden which may aid in Babin M, Cullen JC, Roesler CS, Donaghay PL, Doucette
explaining possible routes of infection and showing possible GJ, Kahru M, Lewis MR, Scholin CA, Sieracki ME, Sosik
effects on fish overall health (Cauyan et al. , 2013). This is in HM, 2005. New approaches and technologies for
addition to taxonomic work on parasitic isopods that have observing harmful algal blooms.
elucidated the presence of Corallana grandiventra in Lake Baldia S, Arguel MKC, Sia D, Tuazon CR, Yap AA, Ong RJ,
Taal and in the country (Tables 1 and 2). Rodis KM, 2011. Comparative analysis of growth and
lipid production in two microalgal species, Botryococcus
CONCLUSIONS AND FUTURE PROSPECTS braunii and Fragilaria brevistriata by nitrogen limitation
experiment. Acta Manilana 59:57-63.
This paper is the first attempt to consolidate the history of Baldia S, Evangelista A, Aralar E, Santiago A, 2007.
freshwater biology research in the Philippines, and shows its Nitrogen and phosphorus utilization in the
major developments and achievements. Furthermore, it cyanobacterium Microcystis aeruginosa isolated from
highlights the growth of the discipline in the University of Laguna de Bay, Philippines. Journal of applied
Santo Tomas - an institution whose alumni played a role phycology 19:607-613.
during the beginnings of the discipline, but was not able to Baldia SF, Conaco MCG, Nishijima T, Imanishi S, Harada KI,
sustain its visibility in the field until the developments of the 2003. Microcystin production during algal bloom
last decade. This was made possible through the synergistic occurrence in Laguna de Bay, the Philippines. Fisheries
approach between key players – faculty researchers, Science 69:110-116.
graduate and undergraduate students, national and Baldia SF, Pantastico JB, Baldia JP. 1986. Efficiency of
international collaborators. This has led to the increased some cyanophytes as larval feed for silver carp
output via peer-reviewed publications, and helped usher in (Hypophthalmichthys molitrix) and the culture of
grants from both national and international funding agencies. Spirulina platensis, p. 609-614. In: Maclean, JL, Dizon,
At present, Thomasian researchers are involved in more LB, Hosillos, LV (eds.). The First Asian Fisheries Forum.
research projects in freshwater biology. It is hoped that this Proceedings of the First Asian Fisheries Forum, 26-31
will be sustained, and would lead to an improvement of May 1986, Manila, Philippines. Manila, Philippines: Asian
scientifically-derived knowledge of freshwater ecosystems in Fisheries Society. pp. 609-614.
the country. Briones JC, Tsai C-H, Nakazawa T, Sakai Y, Papa RDS,
Hsieh C-H, Okuda N, 2012. Long-Term Changes in the
ACKNOWLEDGEMENTS Diet of <italic>Gymnogobius isaza</italic> from Lake
Biwa, Japan: Effects of Body Size and Environmental
We would like to express our gratitude to Acad. Prof. Prey Availability. PloS ONE 7:e53167.
Fortunato B. Sevilla III for suggesting that this paper be made Briones JCA, Papa RDS, Cauyan GA, Mendoza N, Okuda N,
to consolidate a history of UST researches in the field. The 2016. Fish diversity and trophic interactions in Lake
assistance of Elfritzson Peralta and Hazel Guerrero in making Sampaloc (Luzon Is., Philippines). Tropical Ecology 57:in
figures 1 and 3 is also greatly appreciated. This review paper press.
is dedicated to the students who have contributed to the Briones JCA, Papa RDS, Cauyan GA, Urabe M, 2015. The
development of freshwater biology research in the university, first report of three acanthocephalan parasite species
both past and present. isolated from Philippine fishes. . Helminthologia 52:in
press.
LITERATURE CITED Buñag DM, 1934. A supplement to our knowledge of two
Adorador VC, Bernardo MA, Gonzales JLL, Lam AC, Malijan Philippine lakes. Internationale Revue der gesamten
Fam, Papa RDS, Castillo JR, 2006. Morphological and Hydrobiologie und Hydrographie 31:432-435.
DNA size-based differences among Corallana spp. in Cariño JKI, 2003. Integrated water resources management:
Oreochromis sp. found in open waters and fish cages in The experience of Laguna Lake Development Authority
Taal Lake. Philipp. Biota 39:5-11. (LLDA) Philippines. Proceedings of the National
Aquino R, Cho C, Cruz MA, Saguiguit A, Papa RDS, 2008. Congress on Philippine Lakes. Los Baños, Laguna,
Zooplankton composition and diversity in Paoay Lake, Philippines.
Luzon Is., Philippines. Philipp. J. Sci. 137:169-177. Castro SC. 1977. Bioecology of the Freshwater
Arthur JR, Lumanlan-Mayo S. 1997. Checklist of the parasites Prosobranch, Idiopoma angularis (Muller) (Bellamyidae:
of fishes of the Philippines. Technical Paper 369 of the Viviparidae) in Laguna Lake. University of Santo Tomas.
Food and Agriculture Organization of the United Nations Cauyan GA, Briones JCA, De Leon E, Gonong JA,
Fisheries in cooperation with the Fish Health Section of Pasumbal EO, Pelayo MCI, Piñera MA, Papa RDS,
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